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Aspects of the Germination of Some New Zealand Ericaceae

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Aspects of the Germination of Some New Zealand Ericaceae Aspects of the germination of some New Zealand Ericaceae Simon H. Moore1 and Peter Bannister Department of Botany, University of Otago, P.O. Box 56, Dunedin, New Zealand 1Current address: Department of Conservation, Nelson/Marlborough Conservancy, Private Bag 5, Nelson, New Zealand Abstract Seeds of Northern Hemisphere ericoid species are light-sensitive, long-lived, form seed banks, and may be stimulated by short periods of high temperature, but are usually unaffected or show reduced germination in response to low temperature stratification. Comparative study of the germination responses of five New Zealand species of Gaultheria (G. antipoda, G. crassa, G. depressa, G. macrostigma, G. parvula), two hybrids (G. macrostima x crassa, G. macrostigma x depressa) and one introduced European species (Erica lusitanica) showed that light stimulated germination of all species, while germination was not stimulated in dry seed subjected to short periods of high temperature but viability was little affected. Germination of imbibed seed of two other European species (Erica ciliaris, E. erigena) was stimulated by short exposures to high temperature, and germination in E. erigena was inhibited by low temperature stratification. Germination of some New Zealand seed lots was stimulated by low temperature stratification, particularly in collections from high altitude (G. depressa) and in some collections of G. macrostigma and its hybrid with G. depressa. Seeds were largely unaffected by up to ten successive periods of hydration and desiccation, and remained viable after 16 months of dry storage and 12 months of moist storage. There was evidence of after-ripening in seed of G. depressa and G. macrostigma x depressa. Seed lots collected from the same sites in different years showed variable germination responses. Seed germination in Ne"w Zealand Ericaceae is perhaps more similar to Northern Hemisphere Ericaceae than it is to other native woody plants. Additional key words: Erica, Gaultheria, Pemettya, after-ripening, desiccation, heat shock, light/dark, low temperature, stratification low temperature stratification and germination may be Introduction stimulated by short periods of-i exposure to high Ericaceous plants have been widely studied in temperatures (Whittaker and Gimingham, 1962; Ban­ northern and western Europe, largely because of interest nister, 1965; Mallik and GiminghaDi, 1985) and diurnal in heathland, initially as a wasteland that could be made fluctuations in temperature (Gimingham, 1960). more productive and more recently as a disappearing In New Zealand, native ericaceous species are habitat that needs to be preserved (Gimingham, 1992). confined to species of Gaultheria and Pernettya. These Consequently the biology and ecology of British and two genera are differentiated on the basis of fruit European heath species has been well studied and the characters: in Gaultheria a dry capsule is surrounded by germination characteristics of European ericaceous heath a fleshy calyx, whereas in Pernettya the true fruit is a plants are reasonably well known, particularly for ericoid berry with a persistent calyx which may also be species such as heather (Calluna vulgaris) and common succulent. Seeds are numerous and small, and of a species of Erica (E. cinerea, E. tetralix). similar size to those of European ericoid species (Moore, These species have small (< 1 mm in length), long­ 1996). Intergeneric hybrids producing fertile seed occur lived seed that forms persistent seed banks. Afforest­ and, consequently, we have followed Middleton (1991) ation of heathland in Britain leads to the elimination of in this paper and included all Pernettya species within heathland species, such as Calluna vulgaris and Erica Gaultheria. Written information on the germination tetralix, but there is significant regeneration of these requirements of Gaultheria species native to New species when trees are felled after 40 years or more (Hill Zealand appears to be limited to the unpublished work of and Stevens, 1981). As with many species that form Armstrong (1964), a short paper by Bannister (1990) and seed banks, seeds do not germinate in the dark but are brief mentions elsewhere (e.g., Fountain and Outred, stimulated by exposure to light, do not appear to require 1991; Bannister and Jarneson, 1994). Gaultheria species Seed Symposium 1999 83 Germination aspects of some NZ Ericaceae show no or minimal germination in the dark but germin­ seeds were stored either in light-proof canisters (E. ation is stimulated by exposure to light and (in G. lustitanica) or in transparent containers (E. ciliaris, E. macrostigma) may be enhanced by low temperature strat­ erigena) at room temperature. ification (Bannister, 1990; Bannister and Jameson, 1994). Germination of the South American G. mucronata Seed germination appears to respond to low temperature stratification Seeds were plated out on two layers of Whatman No (Arena et al., 1994). The germination ecology of Erica 29 filter paper placed over a layer of absorbent cotton lusitanica, a European ericoid species naturalised in New wool in glass petri dishes. Paper and cotton wool were Zealand, has been investigated by Mather and Williams kept moist with distilled water during the course of the (1990). This paper also presents data for seeds of two experiments. Only apparently filled seeds were used in other European species, Erica erigena and E. ciliaris, the experiments and typically two replicates of 100 seeds collected in the British Isles and germinated in New were used for each treatment. In experiments with E. Zealand. The data for E. ciliaris have been published erigena, only 50 seeds were used in each replicate and elsewhere (Rose et al., 1996). only 25 seeds were used in hydration-dehydration The object of this work was to examine the experiments. germination responses of native Gaultheria species in Seeds of the Gaultheria species and Erica lusitanica more detail and compare them with what is known for were kept in an illuminated cabinet (50 J.tmollm2/s) with Northern Hemisphere Ericaceae. The principal factors a twelve hour alternation of light at 22°C and dark at examined in this paper are responses to high and low l2°C. Petri dishes were laid out in a randomised block temperature, desiccation, after-ripening and longevity. design within the cabinets. Some treatments were started in, or transferred to, an unheated, shaded (c. 56 % of ambient) glasshouse. Seeds of Erica species were ger­ Materials and Methods minated at ambient laboratory temperatures and light. Species and sites of seed collection Radicle emergence was taken as evidence of germination Seed of Erica lusitanica was collected from the and experiments were terminated after about 270 d when Silverpeaks, near Dunedin, in April1992 and April1993, germination had usually ceased, although some experi­ that of E. ciliaris from Dorset, England, in October 1994 ments with Erica erigena were terminated after one year. and that of E. erigena from Errisbeg, Co. Galway, and The final germination percentage was the main statistical Mallaranny and Furnace Lough, Co. Mayo, Ireland, in parameter used in analyses of variance. The inverse of August 1994. Seeds of native New Zealand species were the time taken for 50 % of final germination was also collected from a variety of sites in the southern South used as a function of germination rate, but is referred to Island of New Zealand (site details are given by Moore in the text only if there is a particular point to be made. 1996). In April 1992 and 1993 collections of seed were made from Leith Valley, Dunedin (Gaultheria antipoda) Experimental treatments and the Hawkdun Range, near Undaunted Creek and on Seeds were stratified at 4°C for four weeks, although Otematata Station (G. crassa, G. depressa from 775 m, a 12 week period was also used for Gaultheria 1280 m, 1580 m, G. macrostigma (=Pernettya macro­ macrostigma and G. macrostigma X depressa, and stigma) and G. parvula (=Pernettya nana)). Seeds of G. additional periods of 8 and 12 weeks for E. ciliaris and macrostigma and itshybrids were collected from Flag­ E. erigena. Heat treatments of 30 sec and 2 min at staff, Dunedin, in April 1992 and March 1993 (G. 100°C were used on dry seeds of the Gaultheria species macrostigma, G. macrostigma X depressa, G. macro­ and E. lusitanica, whereas imbibed seeds of the Erica stigma x crassa) with additional collections of G. macro­ ciliaris and E. erigena were soaked in water at 20, 50, stigma in December 1992 and May 1993. and 80°C for five min and at 100°C for one and five Mature fruits were collected from several plants min. within each site. The fleshy fruits of the Gaultheria After-ripening and longevity of seed was tested by species were placed in petri-dishes and allowed to dry in germinating seed stored dry for 1, 3 and 16 months. The the sun. Dried fruits were broken open to release the response of seeds to repeated desiccation and re­ seeds which were separated from any extraneous material hydration was tested for up to ten cycles where seeds and stored in light-proof canisters at room temperature were allowed to imbibe for a week and then desiccated until used. The fruits of the Erica species were already for 24 h before being rehydrated. air-dry on collection and further drying caused their The effects of time of harvest on seed of G. capsules to open and shed their seed. The separated macrostigma collected from Flagstaff at various times Seed Symposium 1999 84 Germination aspects of some NZ Ericaceae during summer and autumn (1992-1993) were also continuous germination through the experimental period. evaluated by both standard germination in the cabinet Further details of germination patterns are given in and by using seed that had been stratified at 4°C for 12 Moore (1996). weeks. Ecotypic comparisons were also made on G. macrostigma from various sites and G. depressa from an Germination in the growth cabinet and in the altitudinal range.
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